Charles Darwin University

Nesting beside old nests, but not over water, increases current nest survival in a tropical mangrove-dwelling warbler

Noske, Richard; Mulyani, Yeni; Lloyd, Penn

Published in: Journal of Ornithology

DOI: 10.1007/s10336-012-0922-3

Published: 01/04/2013

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Citation for published version (APA): Noske, R., Mulyani, Y., & Lloyd, P. (2013). Nesting beside old nests, but not over water, increases current nest survival in a tropical mangrove-dwelling warbler. Journal of Ornithology, 154(2), 517-523. https://doi.org/10.1007/s10336-012-0922-3

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Download date: 24. Sep. 2021 Nesting beside old nests, but not over water, increases current nest survival in a tropical mangrove-dwelling warbler

Richard A. Noske, Yeni A. Mulyani & Penn Lloyd

Journal of Ornithology

ISSN 2193-7192 Volume 154 Number 2

J Ornithol (2013) 154:517-523 DOI 10.1007/s10336-012-0922-3

1 23 Your article is published under the Creative Commons Attribution license which allows users to read, copy, distribute and make derivative works, as long as the author of the original work is cited. You may self- archive this article on your own website, an institutional repository or funder’s repository and make it publicly available immediately.

1 23 J Ornithol (2013) 154:517–523 DOI 10.1007/s10336-012-0922-3

ORIGINAL ARTICLE

Nesting beside old nests, but not over water, increases current nest survival in a tropical mangrove-dwelling warbler

Richard A. Noske • Yeni A. Mulyani • Penn Lloyd

Received: 11 July 2012 / Revised: 25 November 2012 / Accepted: 29 November 2012 / Published online: 30 January 2013 Ó The Author(s) 2013. This article is published with open access at Springerlink.com

Abstract Nest predation rates often depend on how well they signify ‘safe’ locations that have escaped nest preda- a nest is concealed. Many tropical build their nests tion in the past. over water and/or build large pendant nests that closely resemble clumps of vegetation or flood debris, behaviours that are universally assumed to reduce nest predation rates Keywords Nest location Predation Flooding by deterring or confusing nest predators, respectively. Yet Large-billed this hypothesis has never been tested. In tropical Australian mangroves, Large-billed Gerygones magniros- Zusammenfassung tris typically build their pensile nests over tidal channels and often next to their old nests. We monitored the fate of Nisten neben alten Nestern aber nicht u¨ber Wasser nests of 28 pairs and used generalized linear models to erho¨ht den Nesterfolg eines tropischen, Mangroven investigate the importance of nest location to nesting suc- bewohnenden Singvogels cess. Nest failure due to tidal flooding, but not predation, decreased significantly with nest height and distance from Nestpra¨dationsraten ha¨ngen ha¨ufig davon ab, wie gut the tidal channel, suggesting that nesting over water does versteckt das Nest angelegt wird. Viele tropische Vogel- not always reduce nest predation, and, indeed, may incur arten bauen ihre Nester u¨ber Wasser und/oder bauen große, costs through flooding. However, nest predation was sig- ha¨ngende Nester, welche in ihrem Erscheinen Bu¨ndeln aus nificantly more likely at nests that were distant from old Vegetation oder Getreibsel sehr a¨hneln. Es wird allgemein nests than those built close to old nests. Nest predators angenommen, dass dieses Verhalten die Nestpra¨dation might give up searching for eggs or nestlings if nearby verringert, indem Pra¨datoren entweder abgehalten oder nests are empty (potential-prey-site hypothesis). Alterna- irritiert werden. Diese Hypothese wurde bisher jedoch tively, gerygones may build near old intact nests because nicht getestet. In Mangrovenwa¨ldern des tropischen Aust- raliens baut die Sumpfgerygone Gerygone magnirostris ihr ha¨ngendes Nest typischerweise u¨ber Gezeitenkana¨len und ha¨ufig in na¨chster Na¨he zu a¨lteren Nestern. Es wurde der Brutverlauf an den Nestern von 28 Brutpaaren unter- sucht und mit Hilfe linearer Modelle der Einfluss des Communicated by J. Fjeldsa˚. Nistplatzes auf den Erfolg der Brut analysiert. Es zeigte sich eine signifikant geringere Ausfallrate durch U¨ berflu- R. A. Noske (&) Y. A. Mulyani School of Environmental and Life Sciences, tung, nicht aber durch Pra¨dation, je ho¨her das Nest in der Charles Darwin University, Darwin, NT 0909, Australia Vegetation angelegt wurde und mit zunehmender Distanz e-mail: [email protected] des Neststandorts zum Gezeitenkanal. Daher wird ange- nommen, dass die Nestanlage u¨ber Wasser nicht generell P. Lloyd Biodiversity Assessment and Management, Cleveland, zu geringerer Pra¨dation fu¨hrt und ho¨here Risiken durch QLD 4163, Australia U¨ berflutung mit sich bringt. Die Wahrscheinlichkeit der 123 518 J Ornithol (2013) 154:517–523

Nestpra¨dation war jedoch signifikant ho¨her bei Nestern, die vulnerable to flooding by rainwater runoff or high tides in gro¨ßerer Entfernung zu a¨lteren Nestern angelegt waren. (Franklin and Noske 2000; Hudgens 1997; MacGillivray Nestpra¨datoren ko¨nnten die Suche nach Gelegen aufgeben, 1918). Likewise, many tropical species build large pendant wenn nahe gelegene Nester leer sind (,,potential-prey-site nests that closely resemble clumps of vegetation or flotsam hypothesis‘‘). Ein alternativer Erkla¨rungsansatz wa¨re, dass (Bruce 2003; Collias and Collias 1984; Fitzpatrick 2004), die Vo¨gel in der Na¨he alter, intakter Nester bauen, and whilst this resemblance may be viewed as a form of weil diese ,,sichere‘‘Neststandorte ohne vorangegangene camouflage, such nests, particularly when hanging in open Pra¨dationsereignisse andeuten. spaces, are conspicuous to humans and presumably also to visually-oriented nest predators and brood parasites (Biancucci and Martin 2010). Australasian warblers (family ) of the Introduction genus Gerygone are small (6–7 g) insectivorous birds whose nests are typically neat, pendant, oval or dome- Nest predation is the major cause of reproductive failure shaped structures composed of strips of bark, dry grass, among birds (Lack 1968; Martin 1992; Ricklefs 1969), and dead leaves and vines, and featuring a hooded side entrance appears to have played a key role in shaping avian repro- near the top and a small ‘‘tail’’ piece that hangs from the ductive strategies, and hence life history (Fontaine and bottom of the nest. The nest of the tropical Large-billed Martin 2006; Martin 1995; Weidinger 2002). Variation in Gerygone Gerygone magnirostris, on the other hand, is a the location and form of nests has been widely attributed to large and untidy construction, up to 120 cm in length, but avoidance of nest predation (Collias and Collias 1984). averaging around 50 cm long, with the entrance to the nest Nests may be designed to conceal their contents (e.g., chamber in the middle portion, and liberally decorated with enclosed and domed nests) or deceive predators (e.g., dead leaves and other plant materials (Higgins and Peter camouflaged and ‘‘false’’ nests), or may be placed in 2002). These nests are usually suspended conspicuously inaccessible (e.g., cliffs) or well-guarded places (e.g., near from the tips of foliage of trees overhanging rivers in nests of predacious birds or injurious insects) (Collias and mangals (mangrove communities) or rainforests, where Collias 1984). Hole-nesters, especially those that excavate their resemblance to tidal or flood debris has earned the their own nests, suffer less nest predation than those that species the name of ‘‘flood ’’ (Beruldsen 1980; build exposed nests (Martin 1995; Martin and Li 1992; Campbell and Barnard 1917; Le Soue¨f 1903; Marshall Oniki 1979; Skutch 1966; von Haartman 1957; Wesolow- 1933; McGill 1970). This resemblance is tacitly presumed ski and Tomialojc´ 2005). Although well-concealed nest to reduce predation rates by confusing visually-orientated sites might be expected to be safer from nest predators, the nest predators that are unable to distinguish between nests relationship between nest concealment and nest survival is and flotsam. The potential-prey-site hypothesis (Martin confounded by variation in predator faunas and strategies 1993; Martin and Roper 1988) proposes that the probability and parental nest defense behaviour, among other factors of a predator finding stationary prey (such as eggs or (Burhans and Thompson 1998; Colombelli-Ne´grel and nestlings) should decrease with increasing numbers of Kleindorfer 2009;Go¨tmark et al. 1995; Holway 1991; unoccupied potential sites that must be searched, causing Howlett and Stutchbury 1996; Martin and Roper 1988; the predator to give up before finding the occupied site. Nalwanga et al. 2004; Remesˇ 2005; Weidinger 2002). This hypothesis has received recent empirical support However, nest site characteristics other than height and (Chalfoun and Martin 2009). degree of concealment by vegetation are rarely considered As is the case with several other gerygone species, the in studies of nest predation. Large-billed Gerygone often builds its nest close to nests of Many tropical bird species from diverse families (e.g., aggressive colonial (Cornwall 1910; Johnstone Tyrannidae, Eurylaimidae, Ploceidae) are known to build 1990; MacGillivray 1914; Marshall 1933), an association their nests over water (Bruce 2003; Collias and Collias which may reduce nest predation rates by intimidating 1984; Fitzpatrick 2004). Although it is universally accepted potential nest predators (Collias and Collias 1984; Joyce that this behaviour has evolved to hamper access to the nest 1993; Wunderle and Pollock 1985). In addition, active by nest predators (e.g., Collias and Collias 1984; Hudgens nests of this species are sometimes found within 2 m of 1997; Immelmann 1961), and tacitly assumed that such abandoned or old nests, presumably built by the same pair predators are either unable to swim or at least reluctant to (Higgins and Peter 2002), creating a loose cluster of two to enter water, this hypothesis has, to our knowledge, never four nests (Mulyani 2004, unpublished data; Noske, been tested. We propose the name ‘aqua-phobic nest unpublished data). Such disused nests, often with blocked predator’ for this hypothesis. Yet nesting over water may entrances, may act as decoys that confuse predators (Col- incur fitness costs to the owners if such nests become more lias and Collias 1984; Whitney et al. 1996). 123 J Ornithol (2013) 154:517–523 519

In this paper, we examine the relationship between nest Nests were usually monitored every 2–7 days, depend- success and nest site characteristics of Large-billed Gery- ing on their status and accessibility. Nest fate was classified gones in mangals around Darwin, Northern Territory, as successful or failed or unknown. Nest failure was Australia. Artificial nest studies in this region indicated that attributed to one of the following: (1) depredated: with egg nest predation rates are significantly higher in mangals than or nestling missing prior to the expected date of hatching or in other local habitats (Noske et al. 2008), and that within fledging, not after high tide or severe weather, often with mangals, rates are higher in the tall mangrove forest along signs of nest damage; (2) flooded: nests saturated, or dried tidal creeks, which are regularly flooded at high tides, than and stiff, with hood collapsed, usually following a high in the short mangroves surrounding bare, infrequently spring tide; (3) parasitized: nest that successfully produced inundated saltflats (Mulyani 2004; Sato et al. 2010). Little Bronze- (Chrysococcyx minutillus) nestling; We predicted that (1) nests suspended directly over reg- and 4) other causes, which include abandoned nests (nests ularly flooded tidal channels should suffer less nest predation unattended prior to expected date of hatching or fledging, than those further away because non-flying nest predators with cold or cracked eggs or dead nestling) and nests that should avoid the risk of falling into the water (aqua-phobic disappeared, possibly due to drift by tides or storms). nest predator hypothesis); and (2) active nests in close We measured the height of nests less than 2 m above the proximity to old nests should suffer less predation than sol- muddy ground using a 2-m tape measure during a low tide. itary nests, because some predators might give up searching Heights of nests between 2 and 5 m were estimated to the if nearby nests are empty (potential-prey-site hypothesis). nearest 0.2 m, and heights of over 5 m were estimated to We also predicted that low nests would be flooded by very the nearest 0.5 m. Distance to creek was measured using a high tides, as have been observed in the study species else- tape measure. Nest dimensions were measured only after where (MacGillivray 1918), and in an unrelated species in the nests had been depredated or vacated naturally to the study area (Franklin and Noske 2000). minimize disturbance to the breeding birds. For nests that were still in good condition, we measured total nest length, chamber diameter, and entrance width. In addition, we Methods noted the number of old nests within a radius of 10 m from the active nest. Active nests that were built within 5 m of a Two study sites were selected within mangals around Darwin, nest were noted. Due to the nature of working in areas Northern Territory, in northern Australia (12°200–12°280S, that experience large fluctuations in tidal levels, not all 130°480–130°550E; mean annual rainfall, 1,600 mm), where measurements could be taken for all nests. This resulted in the climate is monsoon-tropical, with relatively uniform different sample sizes for some measurements. temperatures year-round, but a very distinct wet season from November to April, when 90 % of the rain fell (Taylor and Data analysis Tulloch 1985). Darwin’s harbour and coastline are macrotidal, with a mean spring range of 5.5 m (maximum, 7.8 m) and We examined relationships between nest fate and depen- mean neap range of 1.9 m (Semenuik 1985). The study sites, dent variables of interest using generalized linear model- Ludmilla Creek (12°230S, 130°510E; and Casuarina Coastal ling (GLM) assuming a binomial error structure (1 = Reserve (12°210S, 130°530E), were situated on and around success, 0 = fail) with a logit link function. We fitted these tidal rivers and pools, with c. 100 % canopy cover of tall models using the software R 2.13.0 (Ihaka and Gentleman White Mangrove Avicennia marina and Stilt Mangrove 1996), following the methods of (Crawley 2002). Model Rhizophora stylosa, and an understory of other shorter man- simplification using backward-elimination of non-signifi- groves, such as Aegiceras corniculatus and Camptostemon cant explanatory variables and interaction terms was schultzii. These sites supported at least 20 pairs and 8 pairs of adopted. Terms were systematically removed from the Large-billed Gerygones, respectively. model and only put back in if their removal resulted in a Intensive nest searches were conducted almost every significant loss of model explanatory power, determined by month from 2000 to early 2003 at both sites. Nests of Large- comparing the log-likelihood of the full model to the log- billed Gerygones are large and conspicuous, being suspended likelihood of the reduced model using the G2 test, whose from the branch tips of mangroves in the open, with no sampling distribution approximates a v2 distribution with vegetation underneath them. Consequently, almost all nests one degree of freedom (Quinn and Keough 2002). The were found by systematically searching along the margins of significance of each explanatory variable was similarly tidal creeks, the preferred habitat of local populations of this determined by comparing the log-likelihood of the full species (Noske 1996). Each nest found was given a unique minimal model including the variable of interest to the code number on blue (1999–2002), red (2001) or pink (2002) log-likelihood of the reduced model excluding it. All flagging tape placed at least 5 m from the nest. meaningful interactions (only two-way interactions) were 123 520 J Ornithol (2013) 154:517–523 included in the saturated model. Because only 42 % of 100 nests had all measurements recorded, if an explanatory variable was found to be non-significant, sample size in the 80 reduced model was increased by including nests that lacked a measurement for that non-significant variable. Residual 60 plots and normal probability plots were used to check for deviations from normality in the final model. To test the 40 prediction that nests located closer to the creek margin and/ No. active nests or at a lower height are at greater risk of mortality from 20 flooding, we undertook a GLM analysis comparing the 0 characteristics of nests that successfully fledged young < 2.0 2.1-4.0 4.1-6.0 6.1-8.0 8.1-10.0 (gerygone and/or cuckoo) and those that failed due to Distance from creek (m) flooding. This analysis excluded nests that failed due to predation, abandonment, nestling death or of unknown fate. Fig. 1 Distance of nests of Large-billed Gerygones Gerygone magnirostris from the bank of the nearest tidal creek (n = 181) To test the combined predictions of the aqua-phobic nest predator hypothesis, that nests located closer to the creek margin and/or at a lower height are at lower risk of mor- wasp nest, whereas 57 nests were situated within 10 m of tality from predation, and the potential-prey-site hypothe- an old nest (i.e. clustered). All nests were located within sis, that nests located within a 10 m radius of old nests are 10 m of the banks of tidal channels (Fig. 1), the mean at lower risk of mortality from predation, we undertook a distance being 2.9 m (SD = 2.8 m; n = 187). The mean GLM analysis comparing the characteristics of nests that height of the nest entrance from the ground, or (for those successfully fledged young (gerygone and/or cuckoo) and nests over water) from the level of the banks of the chan- those that failed due to predation. This analysis included nel, was 1.9 m (SD = 0.7 m, n = 222). the categorical variable DISPERSION with two categories: Nest failure through tidal flooding showed no relation- (1) clustered (nest located within a 10 m radius of an old ship with nest dispersion i.e. whether solitary or close to an gerygone nest); or (2) solitary (nest located further than old nest (v2 = 0.97, P = 0.6), but decreased significantly 10 m from an old gerygone nest). As a further test of the with nest height and distance from the tidal channel nest confusion hypothesis, that nests located closer to an (Tables 1, 2). Nest failure through predation showed no old nest or flood debris are at lower risk of mortality from significant relationship with either distance from the tidal predation, we undertook a similar GLM analysis that channel (v2 = 2.47, P = 0.12) or nest height (v2 = 2.32, included only nests located within a 10 m radius of an old P = 0.13), but predation was significantly more likely at nest. These analyses excluded nests that failed due to solitary nests than at clustered nests (Table 2). Among flooding, abandonment, nestling death or of unknown fate. clustered nests, there was no significant effect of distance to old nests (Table 2). Results Discussion Of 152 known-fate nests, only 21 (14 %) were successful in producing one or more fledglings. Of 131 failed nests, 92 In our study, Large-billed Gerygone nests that were built (70 %) were depredated and 17 (13 %) flooded by high over or close to tidal creeks, or higher in trees, were no tides (Table 1). Only 4 nests were located within 5 m of a more likely to be predated than those further from the creek Table 1 Mean (?SD) height of nests of Large-billed Gerygones or closer to the ground. This finding is inconsistent with the Gerygone magnirostris, and distance of nest to creek bank (m), aqua-phobic nest predator hypothesis, which suggests that according to fate nests over water should be protected from nest predators Nest fate Distance to creek (m) Nest entrance height (m) that are reluctant to enter or risk falling into water, such as some terrestrial mammals or reptiles. The lack of support Flooded 1.95 ± 2.10 (17) 1.67 ± 0.59 (20) for this hypothesis may indicate that the major nest pre- Depredated 3.07 ± 2.68 (92) 1.86 ± 0.54 (105) dators of this species are birds, which can easily access Other causesa 2.53 ± 2.83 (22) 1.99 ± 0.45 (27) nests by flying to them, or mammals and reptiles that are Total failed 2.83 ± 2.58 (131) 1.86 ± 0.54 (152) adept swimmers. Indeed, in artificial nests mimicking those Total success 4.10 ± 3.97 (21) 2.00 ± 0.74 (27) of the Large-billed Gerygone, Sato et al. (2010) found that a Includes nest abandonment and nest disappearance (due to storm or the marks left by ‘predators’ on plasticine eggs in almost unknown causes) all cases of ‘depredation’ were from birds. Moreover the 123 J Ornithol (2013) 154:517–523 521

Table 2 Summary of model Predictor terms v2 P Effect size ± SE terms and statistics for minimal generalised linear models Nest survival (successful vs. flooded, n = 20 successful and 17 flooded nests) examining the relationship between nest fate Constant -4.70 ± 1.76 (successful = 1, fail = 0) and Nest height (m) 8.95 0.003 2.09 ± 0.85 dependent variables of interest Distance to creek (m) 6.91 0.009 0.37 ± 0.17 Nest survival (successful vs. predation, n = 14 successful and 51 depredated nests) Constant -0.65 ± 0.37 Dispersion: solitary 6.44 0.011 -1.66 ± 0.71 Nest survival (clustered nests; 11 successful and 21 depredated) Constant -0.65 ± 0.37 Distance to old nest (m) 3.01 0.08 No interaction terms were Nest height (m) 2.98 0.08 significant only predation events witnessed on real nests of this spe- oriented nest predators, and induce them to stop searching, cies involved the Black Butcherbird Cracticus quoyi then one would expect ‘predation’ of artificial nests to be (Mulyani 2004; Sato et al. 2010). The lack of observations low, once such nests were found to be unproductive. or evidence of predation of Large-billed Gerygone nests by Indeed, the ‘predation’ rate of artificial domed nests with terrestrial (e.g., mammal faeces, severely damaged plasticine eggs was lower than the predation rate of natural nests) casts further doubt on the aqua-phobic nest predator nests in Large-billed Gerygones (Sato et al. 2010) and hypothesis, and suggests that factors other than nest pre- Superb Fairy-wrens Malurus cyaneus (Colombelli-Ne´grel dation are involved in nest site selection of this species. and Kleindorfer 2009), although the difference may have As expected, our data also show that Large-billed been due to the absence of parental activity (Martin et al. Gerygone nests built over or close to tidal creeks, or closer 2000) or other cues at artificial nests, as well as other to the ground, were more likely to be flooded than those factors (Colombelli-Ne´grel and Kleindorfer 2009). further from the creek or higher in trees. Inundation from However, a simple alternative explanation for the lower spring tides each fortnight, especially during periods of predation rate of nests close to old nests is that the latter, heavy rainfall, therefore imposes a fitness cost on nesting particularly if intact, signify locations that have escaped too close to the ground or tidal channel. Placing the nest damage by nest predators. By experimentally adding eggs over water also presumably poses a risk for young when to real nests from the previous year, showed that nest sites they leave the nest as there is no foliage underneath the of four ground-nesting species had strong repeatable effects nest which could prevent them from falling into the water on nest predation rates, with nests in vulnerable sites should they fail to maintain sufficient altitude on their first consistently being quickly depredated. Hass (1998) showed flight. However, this risk would be even greater for the full- that migratory American Robins moved shorter distances grown nestling Little Bronze-cuckoo, which has twice the between nests when re-nesting after successfully raising mass (15 g) of a full-grown Large-billed Gerygone. one brood than did robins that re-laid after natural or Therefore, an alternative hypothesis, but one that we do not experimental nesting failure, suggesting that they selected test, is that building nests over water selects against brood breeding sites based on their previous nesting success. parasitism by the Little Bronze-cuckoo. Within a breeding season, pairs of Spotted Antbirds pre- Possibly the most significant finding of our study is that ferred to reuse previously successful nest sites overall, and active nests situated close to old intact nests suffered less were more likely to return to these sites for a consecutive predation than nests that were on their own. This finding is nest attempt than they were to previously depredated nest consistent with the potential-prey-site hypothesis as nest sites, presumably to avoid sites recently discovered by predators might give up searching for eggs or nestlings if predators (Styrsky 2005). nearby nests are empty or have no opening. Similarly Building new nests close to old one might also have an Galligan and Kleindorfer (2008) argued that in the Yellow- anti-predator function for incubating female gerygones, rumped Thornbill Acanthiza chrysorrhoa, which builds a which are susceptible to predation as their enclosed nests cup-shaped ‘‘false nest’’ on top of its real domed nest, offer little chance of escape if the predator is able to reach conspicuous nests had higher nesting success than less well the entrance. Predators landing on or climbing to old nests concealed nests because nest predators choose to conserve might cause sufficient movement of the supporting branch energy and time by not examining the nest site more to warn the incubating bird in the active nest of impending closely. If such ‘decoy’ nests served to distract visually- danger.

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Although there are many reports in the literature of Chalfoun AD, Martin TE (2009) Habitat structure mediates predation Large-billed Gerygones nesting near wasp nests, we found risk for sedentary prey: experimental tests of alternative hypotheses. J Anim Ecol 78:497–503 few of the latter near gerygone nests at our study sites. Yet Collias NE, Collias EC (1984) Nest building and bird behaviour. there are no published reports of this species nesting close Princeton University Press, Princeton to old nests, and only two of 19 nest records in the Birds Colombelli-Ne´grel D, Kleindorfer S (2009) Nest height, nest Australia Nest Record Scheme were reported as being built concealment, and predator type predict nest predation in superb fairy-wrens (Malurus cyaneus). Ecol Res 24:921–928 \1.5 m from old or inactive nests, while in two other cases Cornwall EM (1910) Notes on the Great-billed Heron (Ardea where two nests were separated by only 50 cm, both nests sumatrana). Emu 9:138–141 reputedly contained eggs or young (Higgins and Peter Crawley MJ (2002) Statistical computing: an introduction to data 2002). The elevated frequency of clustered nests in our analysis using S-Plus. Wiley, Chichester Fitzpatrick JW (2004) Family Tyrannidae (Tyrant-flycatchers). In: del study area may thus be related to lower abundance of Hoyo J, Elliott A, Christie DA (eds) Handbook of the birds of the wasps, or to elevated nest predation and/or brood parasit- world. vol 9 Cotingas to Pipits and Wagtails. Lynx, Barcelona, ism levels. Nest predation rates during the study period pp 170–462 were not exceptionally high (57 %; n = 160) in this spe- Fontaine JJ, Martin TE (2006) Parent birds assess nest predation risk and adjust their reproductive strategies. Ecol Lett 9:428–434 cies, whereas an astonishing 41 % of nests were parasitised Franklin DC, Noske RA (2000) The nesting biology of the Brown by the Little Bronze-cuckoo, although many of these nests Lichmera indistincta in the Darwin Region of were also depredated (Mulyani 2004). Brood parasitism northern Australia with notes on tidal flooding of nests. Corella rates were also high (34 %; n = 138) for the coexisting 20:38–44 Galligan TH, Kleindorfer S (2008) Support for the nest mimicry Mangrove Gerygone G. levigaster, whose nests were never hypothesis in Yellow-rumped Thornbills Acanthiza chrysorrhoa. observed close to older nests, but this species builds its Ibis 150:550–557 nests within the foliage of small mangrove shrubs on bare Go¨tmark F, Blomqvist D, Johansson OC, Bergqvist J (1995) Nest-site saltflats rather than in the open space over tidal rivers selection: a trade-off between concealment and view of the surroundings. J Avian Biol 26:305–312 (Mulyani 2004; Noske 2001). Brood parasitism can cause Hass CA (1998) Effects of prior nesting success on site fidelity and higher reproductive costs than nest predation due to delays breeding dispersal: an experimental approach. Auk 115:929–936 in re-nesting resulting from time expended in foster care, Higgins PJ, Peter JM (eds) (2002) Handbook of Australian, New and thus may constitute a stronger selection pressure on Zealand and Antarctic birds. vol 6 Pardalotes to shrike-thrushes. Oxford University Press, Melbourne these species than nest predation in the study area. Holway DA (1991) Nest-site selection and the importance of nest concealment in the Black-throated Blue Warbler. Condor Acknowledgments We thank C. Brady, S. Manullang, Y. Widi- 95:83–93 wardono, R. Funnel, V. Vilar and numerous volunteers who helped in Howlett JS, Stutchbury BJ (1996) Nest concealment and predation in locating nests. We also received substantial help in statistical analysis Hooded Warblers: experimental removal of nest cover. Auk from K. McGuiness, D. Franklin, and W. Telfer. This study was a part 113:1–10 of PhD research of the second author under the AusAID scholarship Hudgens BR (1997) Nest predation avoidance by the Blue-billed scheme. Malimbe Malimbus nitens (Ploceinae). Ibis 139:692–694 Ihaka R, Gentleman R (1996) R: a language for data analysis and Open Access This article is distributed under the terms of the graphics. J Comput Graph Stat 5:299–314 Creative Commons Attribution License which permits any use, dis- Immelmann K (1961) A contribution to the biology and ethology of tribution, and reproduction in any medium, provided the original Australian (Meliphagidae). Z Ornithol 102:164–207 author(s) and the source are credited. Johnstone RE (1990) Mangroves and mangrove birds of Western Australia. Western Australian Museum, Perth Joyce FL (1993) Nesting success of rufous-naped wrens (Camp- ylorhynchus rufinucha) is greater near wasp nests. Behav Ecol Sociobiol 32:71–77 References Lack D (1968) Ecological adaptations for breeding in birds. Methuen, London Beruldsen G (1980) A field guide to nest and eggs of Australian birds. Le Soue¨f WHD (1903) Description of birds’ eggs from Port Darwin Rigby, Adelaide District, Northern Australia. Emu 2:139–159 Biancucci L, Martin TE (2010) Can selection on nest size from nest MacGillivray W (1914) Notes on some North Queensland birds. Emu predation explain the latitudinal gradient in clutch size? J Anim 13:132–186 Ecol 79:1086–1092 MacGillivray W (1918) Ornithologists in North Queensland. Emu Bruce M (2003) Family Eurylaimidae (Broadbills). In: del Hoyo J, 17:180–212 Elliott A, Christie DA (eds) Handbook of the birds of the world, Marshall AJ (1933) The Large-billed warbler. Emu 33:81–85 vol 8: Broadbills to Tapaculos. Lynx, Barcelona, pp 54–93 Martin TE (1992) Interaction of nest predation and food limitation in Burhans DE, Thompson FR III (1998) Effects of time and nest-site reproductive strategies. Curr Ornithol 9:163–197 characteristics on concealment of songbird nests. Condor Martin TE (1993) Nest predation and nest sites. Bioscience 43: 100:663–672 523–532 Campbell AJ, Barnard HG (1917) Birds of the Rockingham Bay Martin TE (1995) Avian life history evolution in relation to nest sites, District, North Queensland. Emu 17:2–38 nest predation and food. Ecol Monogr 65:101–127

123 J Ornithol (2013) 154:517–523 523

Martin TE, Li P (1992) Life history traits of open- vs cavity-nesting Ricklefs RE (1969) An analysis of nesting mortality in birds. birds. Ecology 73:579–592 Smithson Contrib Zool 9:1–48 Martin TE, Roper JJ (1988) Nest predation and nest site selection in a Sato NJ, Morimoto G, Noske RA, Ueda K (2010) Nest form, colour, western population of the hermit thrush. Condor 90:51–57 and nesting habitat affect predation rates of Australasian Martin TE, Scott J, Menge C (2000) Nest predation increases with warblers (Gerygone spp.) in tropical mangroves. J Yamashina parental activity: separating nest site and parental activity effects. Inst Ornithol 42:65–78 Proceedings of the Royal Society of London B 267:2287–2293 Semenuik V (1985) Mangrove environments of Port Darwin, McGill AR (1970) Australian warblers. Birds Observers Club, Northern Territory: the physical framework and habitats. J R Melbourne Soc WA 67:81–97 Mulyani YA (2004) Reproductive ecology of tropical mangrove-dwelling Skutch AF (1966) A breeding bird census and nesting success in warblers: the roles of nest predation, brood parasitism and food Central America. Ibis 108:1–16 limitation. PhD thesis, Charles Darwin University, Darwin Styrsky JN (2005) Influence of predation on nest-site reuse by an Nalwanga D, Lloyd P, Du Plessis MA, Martin TE (2004) The open-cup nesting neotropical . Condor 107:133–137 influence of nest-site characteristics on the nesting success of the Taylor JA, Tulloch D (1985) Rainfall in the wet-dry tropics: extreme Karoo Prinia (Prinia maculosa). Ostrich 75:269–274 events at Darwin and similarities between years during period, Noske RA (1996) Abundance, zonation, and foraging ecology of birds 1870–1983 inclusive. Aust J Ecol 10:281–295 in mangroves of Darwin Harbour, Northern Territory. Wildl Res Von Haartman L (1957) Adaptation in hole-nesting birds. Evolution 23:443–474 11:294–347 Noske RA (2001) The breeding biology of the mangrove gerygone, Weidinger K (2002) Interactive effects of concealment, parental Gerygone laevigaster, in the Darwin region, with notes on brood behaviour and predators on the survival of open passerine nests. parasitism by the little bronze-cuckoo, Chrysococcyx minutillus. J Anim Ecol 71:424–437 Emu 101:129–135 Wesolowski T, Tomialojc´ L (2005) Nest sites, nest depredation, and Noske RA, Fischer S, Brook BW (2008) Artificial nest predation rates productivity of avian broods in a primeval temperate forest: do vary among habitats in the Australian monsoon tropics. Ecol Res the generalisations hold? J Avian Biol 36:361–367 23:519–527 Whitney BM, Pacheco JF, Da Fonseca PSM, Barth RH Jr (1996) The Oniki Y (1979) Is nesting success of birds low in the tropics? nest and nesting ecology of Acrobatornis fonsecai (Furnariidae), Biotropica 11:60–69 with implications for intrafamilial relationships. Wilson Bull Quinn GP, Keough MJ (2002) Experimental design and data analysis 108:434–448 for biologists. Cambridge University Press, Melbourne Wunderle JM Jr, Pollock KH (1985) The Banaquit-wasp nesting Remesˇ V (2005) Birds and rodents destroy different nests: a study of association: a random choice model. Ornithol Monogr 36: blackcap Sylvia atricapilla using the removal of nest conceal- 595–603 ment. Ibis 147:213–216

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